But it is a most remarkable fact, that, while the groups which have been mentioned, and many besides, exhibit no sign of progressive modification, there are others, coexisting with them, under the same conditions, in which more or less distinct indications of such a process seem to be traceable. Among such indications I may remind you of the predominance of Holostome Gasteropoda in the older rocks as compared with that of Siphonostome Gasteropoda in the later. A case less open to the objection of negative evidence, however, is that afforded by the Tetrabranchiate Cephalopoda, the forms of the shells and of the septal sutures exhibiting a certain increase of complexity in the newer genera. Here, however, one is met at once with the occurrence of Orthoceras and Baculites at the two ends of the series, and of the fact that one of the simplest genera, Nautilus, is that which now exists.
The Crinoidea, in the abundance of stalked forms in the ancient formations as compared with their present rarity, seem to present us with a fair case of modification from a more embryonic towards a less embryonic condition. But then, on careful consideration of the facts, the objection arises that the stalk, calyx, and arms of the palæozoic Crinoid are exceedingly different from the corresponding organs of a larval Comatula; and it might with perfect justice be argued that Actinocrinus and Eucalyptocrinus, for example, depart to the full as widely, in one direction, from the stalked embryo of Comatula, as Comatula itself does in the other.
The Echinidea, again, are frequently quoted as exhibiting a gradual passage from a more generalized to a more specialized type, seeing that the elongated, or oval, Spatangoids appear after the spheroidal Echinoids. But here it might be argued, on the other hand, that the spheroidal Echinoids, in reality, depart further from the general plan and from the embryonic form than the elongated Spatangoids do; and that the peculiar dental apparatus and the pedicellariæ of the former are marks of at least as great differentiation as the petaloid ambulacra and semitæ of the latter.
Once more, the prevalence of Macrurous before Brachyurous Podophthalmia is, apparently, a fair piece of evidence in favour of progressive modification in the same order of Crustacea; and yet the case will not stand much sifting, seeing that the Macrurous Podophthalmia depart as far in one direction from the common type of Podophthalmia, or from any embryonic condition of the Brachyura, as the Brachyura do in the other; and that the middle terms between Macrura and Brachyura—the Anomura—are little better represented in the older Mesozoic rocks than the Brachyura are.
None of the cases of progressive modification which are cited from among the Invertebrata appear to me to have a foundation less open to criticism than these; and if this be so, no careful reasoner would, I think, be inclined to lay very great stress upon them. Among the Vertebrata, however, there are a few examples which appear to be far less open to objection.
It is, in fact, true of several groups of Vertebrata which have lived through a considerable range of time, that the endoskeleton (more particularly the spinal column) of the older genera presents a less ossified, and, so far, less differentiated, condition than that of the younger genera. Thus the Devonian Ganoids, though almost all members of the same sub-order as Polypterus, and presenting numerous important resemblances to the existing genus, which possesses biconcave vertebræ, are, for the most part, wholly devoid of ossified vertebral centra. The Mesozoic Lepidosteidæ, again, have, at most, biconcave vertebræ, while the existing Lepidosteus has Salamandroid, opisthocœlous, vertebræ. So, none of the Palæozoic Sharks have shown themselves to be possessed of ossified vertebræ, while the majority of modern Sharks possess such vertebræ. Again, the more ancient Crocodilia and Lacertilia have vertebræ with the articular facets of their centra flattened or biconcave, while the modern members of the same group have them procœlous. But the most remarkable examples of progressive modification of the vertebral column, in correspondence with geological age, are those afforded by the Pycnodonts among fish, and the Labyrinthodonts among Amphibia.
The late able ichthyologist Heckel pointed out the fact, that, while the Pycnodonts never possess true vertebral centra, they differ in the degree of expansion and extension of the ends of the bony arches of the vertebræ upon the sheath of the notochord; the Carboniferous forms exhibiting hardly any such expansion, while the Mesozoic genera present a greater and greater development, until, in the Tertiary forms, the expanded ends become suturally united so as to form a sort of false vertebra. Hermann von Meyer, again, to whose luminous researches we are indebted for our present large knowledge of the organization of the older Labyrinthodonts, has proved that the Carboniferous Archegosaurus had very imperfectly developed vertebral centra, while the Triassic Mastodonsaurus had the same parts completely ossified.[38]
The regularity and evenness of the dentition of the Anoplotherium, as contrasted with that of existing Artiodactyles, and the assumed nearer approach of the dentition of certain ancient Carnivores to the typical arrangement, have also been cited as exemplifications of a law of progressive development, but I know of no other cases based on positive evidence which are worthy of particular notice.
What then does an impartial survey of the positively ascertained truths of palæontology testify in relation to the common doctrines of progressive modification, which suppose that modification to have taken place by a necessary progress from more to less embryonic forms, or from more to less generalized types, within the limits of the period represented by the fossiliferous rocks?
It negatives those doctrines; for it either shows us no evidence of any such modification, or demonstrates it to have been very slight; and as to the nature of that modification, it yields no evidence whatsoever that the earlier members of any long-continued group were more generalized in structure than the later ones. To a certain extent, indeed, it may be said that imperfect ossification of the vertebral column is an embryonic character; but, on the other hand, it would be extremely incorrect to suppose that the vertebral columns of the older Vertebrata are in any sense embryonic in their whole structure.